Drought-deciduous Trees Research Articles

Root and branch hydraulic functioning and trait coordination across organs in drought-deciduous and evergreen tree species of a subtropical highland forest.

Vessel traits are key in understanding trees' hydraulic efficiency, and related characteristics like growth performance and drought tolerance. While most plant hydraulic studies have focused on aboveground organs, our understanding of root hydraulic functioning and trait coordination across organs remains limited. Furthermore, studies from seasonally dry (sub-)tropical ecosystems and mountain forests are virtually lacking and uncertainties remain regarding potentially different hydraulic strategies of plants differing in leaf habit. Here, we compared wood anatomical traits and specific hydraulic conductivities between coarse roots and small branches of five drought-deciduous and eight evergreen angiosperm tree species in a seasonally dry subtropical Afromontane forest in Ethiopia. We hypothesized that largest vessels and highest hydraulic conductivities are found in roots, with greater vessel tapering between roots and equally-sized branches in evergreen angiosperms due to their drought-tolerating strategy. We further hypothesized that the hydraulic efficiencies of root and branches cannot be predicted from wood density, but that wood densities across organs are generally related. Root-to-branch ratios of conduit diameters varied between 0.8 and 2.8, indicating considerable differences in tapering from coarse roots to small branches. While deciduous trees showed larger branch xylem vessels compared to evergreen angiosperms, root-to-branch ratios were highly variable within both leaf habit types, and evergreen species did not show a more pronounced degree of tapering. Empirically determined hydraulic conductivity and corresponding root-to-branch ratios were similar between both leaf habit types. Wood density of angiosperm roots was negatively related to hydraulic efficiency and vessel dimensions; weaker relationships were found in branches. Wood density of small branches was neither related to stem nor coarse root wood densities. We conclude that in seasonally dry subtropical forests, similar-sized coarse roots hold larger xylem vessels than small branches, but the degree of tapering from roots to branches is highly variable. Our results indicate that leaf habit does not necessarily influence the relationship between coarse root and branch hydraulic traits. However, larger conduits in branches and a low carbon investment in less dense wood may be a prerequisite for high growth rates of drought-deciduous trees during their shortened growing season. The correlation of stem and root wood densities with root hydraulic traits but not branch wood points toward large trade-offs in branch xylem towards mechanical properties.

Leaf turgor loss point shapes local and regional distributions of evergreen but not deciduous tropical trees.

Summary The effects of climate change on tropical forests will depend on how diverse tropical tree species respond to drought. Current distributions of evergreen and deciduous tree species across local and regional moisture gradients reflect their ability to tolerate drought stress, and might be explained by functional traits.We measured leaf water potential at turgor loss (i.e. ‘wilting point’; πtlp), wood density (WD) and leaf mass per area (LMA) on 50 of the most abundant tree species in central Panama. We then tested their ability to explain distributions of evergreen and deciduous species within a 50 ha plot on Barro Colorado Island and across a 70 km rainfall gradient spanning the Isthmus of Panama.Among evergreen trees, species with lower πtlp were associated with drier habitats, with πtlp explaining 28% and 32% of habitat association on local and regional scales, respectively, greatly exceeding the predictive power of WD and LMA. In contrast, πtlp did not predict habitat associations among deciduous species.Across spatial scales, πtlp is a useful indicator of habitat preference for tropical tree species that retain their leaves during periods of water stress, and holds the potential to predict vegetation responses to climate change.

Phytochemical Investigation and Characterization on the Stem Bark Extract of Croton macrostachyus

Croton macrostachyus which is called ‘rush foil’ or ‘broadleaved croton is a multipurpose, medium sized, drought-deciduous pioneer tree. It is a tall tree found in tropical regions of Africa. The genus Croton belongs to the family Euphorbiaceae, which commonly known as the ‘spurge’ family, and it is known as ‘Bisana’ (in Amharic). Traditionally, C. macrostachyus used for treatment of malaria, rabies, gonorrhea, wound, diarrhea, hepatitis, jaundice, scabies, toothache, abdominal pain, cancer, typhoid, pneumonia and gastrointestinal disorders and as ethno-veterinary medicine. The air dried and powdered plant material (400g) was first soaked with 500mL n-hexane for 48hours and yielded 2g of n-hexane extract. Marc was soaked with 500mL of chloroform for 36hours and afforded 3.5g of chloroform extract. Finally, Marc was soaked with 500mL of methanol and yielded 18g of methanol extract. The chloroform extract of the stem bark of C. macrostachyus afforded a compound coded as EO . Its Structural determination was accomplished by means of spectroscopic techniques, namely IR, 1 H NMR, 13 C NMR and DEPT-135. The compound, EO was isolated and characterized from the stem bark of C. macrostachyus . Generally, more advanced chromatographic techniques are required to isolate more compounds from stem bark of C. macrostachyus . Also MS and 2D NMR spectroscopic techniques are needed to fully characterize the isolated compound. Keywords: C. macrostachyus; ethno-veterinary; characterization; chromatographic techniques; spectroscopic techniques. DOI: 10.7176/CPER/63-01 Publication date: October 31 st 2020

Divergent Hydraulic Safety Strategies in Three Co-occurring Anacardiaceae Tree Species in a Chinese Savanna.

Vulnerability segmentation, the condition under which plant leaves are more vulnerable to drought-induced cavitation than stems, may act as a “safety valve” to protect stems from hydraulic failure. Evergreen, winter-deciduous, and drought-deciduous tree species co-occur in tropical savannas, but there have been no direct studies on the role of vulnerability segmentation and stomatal regulation in maintaining hydraulic safety in trees with these three leaf phenologies. To this end, we selected three Anacardiaceae tree species co-occurring in a Chinese savanna, evergreen Pistacia weinmanniifolia, drought-deciduous Terminthia paniculata, and winter-deciduous Lannea coromandelica, to study inter-species differentiation in leaf and stem hydraulic safety. We found that the two deciduous species had significantly higher sapwood-specific hydraulic conductivity and leaf-specific hydraulic conductance than the evergreen species. Moreover, two deciduous species were more vulnerable to stem cavitation than the evergreen species, although both drought-deciduous species and evergreen species had drought-resistance leaves. The evergreen species maintained a wide hydraulic safety margin (HSM) in stems and leaves; which was achieved by embolism resistance of both stems and leaves and isohydric stomatal control. Both deciduous species had limited HSMs in stems and leaves, being isohydric in the winter-deciduous species and anisohydric in drought-deciduous species. The difference in water potential at 50% loss of hydraulic conductivity between the leaves and the terminal stems (P50leaf−stem) was positive in P. weinmanniifolia and L. coromandelica, whereas, T. paniculata exhibited a lack of vulnerability segmentation. In addition, differences in hydraulic architecture were found to be closely related to other structural traits, i.e., leaf mass per area, wood density, and sapwood anatomy. Overall, the winter-deciduous species exhibits a drought-avoidance strategy that maintains the hydraulic safety of the more carbon-costly stems by sacrificing cheaper and more vulnerable leaves, while the evergreen species exhibits a hydraulic strategy of drought tolerance with strong stomatal regulation. In contrast, the drought-deciduous species lacks vulnerability segmentation and sheds leaves at the expense of top shoots during peak drought. This study demonstrates that even sympatric tree species that differ in leaf phenology can exhibit divergent adaptive hydraulic safety strategies.

Host tree phenology affects vascular epiphytes at the physiological, demographic and community level.

The processes that govern diverse tropical plant communities have rarely been studied in life forms other than trees. Structurally dependent vascular epiphytes, a major part of tropical biodiversity, grow in a three-dimensional matrix defined by their hosts, but trees differ in their architecture, bark structure/chemistry and leaf phenology. We hypothesized that the resulting seasonal differences in microclimatic conditions in evergreen vs. deciduous trees would affect epiphytes at different levels, from organ physiology to community structure. We studied the influence of tree leaf phenology on vascular epiphytes on the Island of Barro Colorado, Panama. Five tree species were selected, which were deciduous, semi-deciduous or evergreen. The crowns of drought-deciduous trees, characterized by sunnier and drier microclimates, hosted fewer individuals and less diverse epiphyte assemblages. Differences were also observed at a functional level, e.g. epiphyte assemblages in deciduous trees had larger proportions of Crassulacean acid metabolism species and individuals. At the population level a drier microclimate was associated with lower individual growth and survival in a xerophytic fern. Some species also showed, as expected, lower specific leaf area and higher δ(13)C values when growing in deciduous trees compared with evergreen trees. As hypothesized, host tree leaf phenology influences vascular epiphytes at different levels. Our results suggest a cascading effect of tree composition and associated differences in tree phenology on the diversity and functioning of epiphyte communities in tropical lowland forests.

Photoprotection of evergreen and drought-deciduous tree leaves to overcome the dry season in monsoonal tropical dry forests in Thailand

In tropical dry forests, uppermost-canopy leaves of evergreen trees possess the ability to use water more conservatively compared with drought-deciduous trees, which may result from significant differences in the photoprotective mechanisms between functional types. We examined the seasonal variations in leaf gas exchange, chlorophyll fluorescence and the amounts of photosynthetic pigments within lamina of the uppermost-canopy leaves of three drought-deciduous trees (Vitex peduncularis Wall., Xylia xylocarpa (Roxb.) W. Theob., Shorea siamensis Miq.), a semi-deciduous tree (Irvingia malayana Miq.) and two evergreen trees (Hopea ferrea Lanessan and Syzygium cumini (L.) Skeels) in Thailand. Area-based maximum carbon assimilation rates (Amax) decreased during the dry season, except in S. siamensis. The electron transport rate (ETR) remained unchanged in deciduous trees, but decreased during the dry season in evergreen and semi-deciduous trees. In the principal component analysis, the first axis (Axis 1) accounted for 44.3% of the total variation and distinguished deciduous from evergreen trees. Along Axis 1, evergreen trees were characterized by a high Stern-Volmer non-photochemical quenching coefficient (NPQ), high xanthophyll cycle pigments/chlorophyll and a high de-epoxidation state of the xanthophyll cycle, whereas the deciduous trees were characterized by a high ETR, a high quantum yield of PSII (ΦPSII = (Fm(') -F)/Fm(')) and a high mass-based Amax under high-light conditions. These findings indicate that drought-deciduous trees showing less conservative water use tend to dissipate a large proportion of electron flow through photosynthesis or alternative pathways. In contrast, the evergreens showed more conservative water use, reduced Amax and ETR and enhanced NPQ and xanthophyll cycle pigments/chlorophyll during the dry season, indicating that down-regulated photosynthesis with enhanced thermal dissipation of excess light energy played an important role in photoprotection. Trees with different water uses and leaf lifespans appear to employ different photoprotective mechanisms to overcome the unfavorable dry-season drought. Our data may suggest that future changes in precipitation will strongly impinge on forest structure and functions.

Droughts, hydraulic redistribution, and their impact on vegetation composition in the Amazon forest

Hydraulic redistribution (HR), the nocturnal transport of moisture by plant roots from wetter to drier portions of the root zone, in general can buffer plants against seasonal water deficits. However, its role in longer droughts and its long-term ecological impact are not well understood. Based on numerical model experiments for the Amazon forest, this modeling study indicates that the impact of HR on plant growth differs between droughts of different time scales. While HR increases transpiration and plant growth during regular dry seasons, it reduces dry season transpiration and net primary productivity (NPP) under extreme droughts such as those during El Nino years in the Amazon forest. This occurs because, in places where soil water storage is not able to sustain the ecosystem through the dry season, the HR-induced acceleration of moisture depletion in the early stage of the dry season reduces water availability for the rest of the dry season and causes soil moisture to reach the wilting point earlier. This gets exacerbated during extreme droughts, which jeopardizes the growth of trees that are not in dry season dormancy, i.e., evergreen trees. As a result, the combination of drought and HR increases the percentage of drought deciduous trees at the expense of evergreen trees, and the fractional coverage of forest canopy is characterized by sudden drops following extreme droughts and slow recovery afterwards. The shift of the tropical forest towards more drought deciduous trees as a result of the combined effects of extreme drought and HR has important implications for how vegetation will respond to future climate changes.

Water relations of evergreen and drought-deciduous trees along a seasonally dry tropical forest chronosequence

Seasonally dry tropical forests (SDTF) are characterized by pronounced seasonality in rainfall, and as a result trees in these forests must endure seasonal variation in soil water availability. Furthermore, SDTF on the northern Yucatan Peninsula, Mexico, have a legacy of disturbances, thereby creating a patchy mosaic of different seral stages undergoing secondary succession. We examined the water status of six canopy tree species, representing contrasting leaf phenology (evergreen vs. drought-deciduous) at three seral stages along a fire chronosequence in order to better understand strategies that trees use to overcome seasonal water limitations. The early-seral forest was characterized by high soil water evaporation and low soil moisture, and consequently early-seral trees exhibited lower midday bulk leaf water potentials (ΨL) relative to late-seral trees (−1.01 ± 0.14 and −0.54 ± 0.07 MPa, respectively). Although ΨL did not differ between evergreen and drought-deciduous trees, results from stable isotope analyses indicated different strategies to overcome seasonal water limitations. Differences were especially pronounced in the early-seral stage where evergreen trees had significantly lower xylem water δ18O values relative to drought-deciduous trees (−2.6 ± 0.5 and 0.3 ± 0.6‰, respectively), indicating evergreen species used deeper sources of water. In contrast, drought-deciduous trees showed greater enrichment of foliar 18O (∆18Ol) and 13C, suggesting lower stomatal conductance and greater water-use efficiency. Thus, the rapid development of deep roots appears to be an important strategy enabling evergreen species to overcome seasonal water limitation, whereas, in addition to losing a portion of their leaves, drought-deciduous trees minimize water loss from remaining leaves during the dry season.

Seasonal variations of gas exchange and water relations in deciduous and evergreen trees in monsoonal dry forests of Thailand

This study compared leaf gas exchange, leaf hydraulic conductance, twig hydraulic conductivity and leaf osmotic potential at full turgor between two drought-deciduous trees, Vitex peduncularis Wall. and Xylia xylocarpa (Roxb.) W. Theob., and two evergreen trees, Hopea ferrea Lanessan and Syzygium cumini (L.) Skeels, at the uppermost canopies in tropical dry forests in Thailand. The aims were to examine (i) whether leaf and twig hydraulic properties differ in relation to leaf phenology and (ii) whether xylem cavitation is a determinant of leaf shedding during the dry season. The variations in almost all hydraulic traits were more dependent on species than on leaf phenology. Evergreen Hopea exhibited the lowest leaf-area-specific twig hydraulic conductivity (leaf-area-specific K(twig)), lamina hydraulic conductance (K(lamina)) and leaf osmotic potential at full turgor (Ψ(o)) among species, whereas evergreen Syzygium exhibited the highest leaf-area-specific K(twig), K(lamina) and Ψ(o). Deciduous Xylia had the highest sapwood-area-specific K(twig), along with the lowest Huber value (sapwood area/leaf area). More negative osmotic Ψ(o) and leaf osmotic adjustment during the dry season were found in deciduous Vitex and evergreen Hopea, accompanied by low sapwood-area-specific K(twig). Regarding seasonal changes in hydraulics, no remarkable decrease in K(lamina) and K(twig) was found during the dry season in any species. Results suggest that leaf shedding during the dry season is not always associated with extensive xylem cavitation.

Role of dynamic vegetation in regional climate predictions over western Africa

This study examines the role of vegetation dynamics in regional predictions of future climate change in western Africa using a dynamic vegetation model asynchronously coupled to a regional climate model. Two experiments, one for present day and one for future, are conducted with the linked regional climate-vegetation model, and the third with the regional climate model standing alone that predicts future climate based on present-day vegetation. These simulations are so designed in order to tease out the impact of structural vegetation feedback on simulated climate and hydrological processes. According to future predictions by the regional climate-vegetation model, increase in LAI is widespread, with significant shift in vegetation type. Over the Guinean Coast in 2084–2093, evergreen tree coverage decreases by 49% compared to 1984–1993, while drought deciduous tree coverage increases by 56%. Over the Sahel region in the same period, grass cover increases by 31%. Such vegetation changes are accompanied by a decrease of JJA rainfall by 2% over the Guinean Coast and an increase by 23% over the Sahel. This rather small decrease or large increase of precipitation is largely attributable to the role of vegetation feedback. Without the feedback effect from vegetation, the regional climate model would have predicted a 5% decrease of JJA rainfall in both the Guinean Coast and the Sahel as a result of the radiative and physiological effects of higher atmospheric CO2 concentration. These results demonstrate that climate- and CO2-induced changes in vegetation structure modify hydrological processes and climate at magnitudes comparable to or even higher than the radiative and physiological effects, thus evincing the importance of including vegetation feedback in future climate predictions.

Potential future changes of the terrestrial ecosystem based on climate projections by eight general circulation models

A number of previous modeling studies have assessed the implications of projected CO2‐induced climate change for future terrestrial ecosystems. However, although current understanding of possible long‐term response of vegetation to elevated CO2 and CO2‐induced climate change in some geographical areas (e.g., the high‐latitude regions) has been strengthened by dint of accumulating evidence from these past studies, it is still weak in others. This study examines the responses of global potential natural vegetation distribution, net primary production (NPP), and fire emissions to future changes in atmospheric CO2 concentration and climate using the National Center for Atmospheric Research Community Land Model's dynamic global vegetation model. The model is run to vegetative equilibrium (i.e., with respect to leaf area index (LAI) and vegetation coverage) driven with preindustrial climate and future climate near 2100, respectively, simulated by eight general circulation models (GCMs). The simulated potential vegetation under the preindustrial control mean climate (CO2 concentration held at 275 ppm) is compared with that under the SRESA1B 2100 mean climate (CO2 concentration stabilizes at 720 ppm beyond 2100). Simulated vegetation response ranges from mild changes of the fractional coverage of different plant functional types to the rather dramatic changes of dominant plant functional types. Although such response differs significantly across different GCM climate projections, a quite consistent spatial pattern emerges, characterized by a considerable poleward spread or shift of temperate and boreal forests in the Northern Hemisphere high latitudes, and a substantial degradation of vegetation type in the tropics (e.g., increase of drought deciduous trees coverage at the expense of evergreen trees) especially in portions of West and southern Africa and South America. Despite the widespread degradation of vegetation type in the tropics, NPP, and growing season LAI are predicted to increase under most GCM scenarios over most of the globe. Carbon fluxes to the atmosphere due to fire generally increase too across the globe. Such responses of NPP and fire occurrence result from the synergistic effects of CO2 concentration changes, climate changes, and vegetation changes. In the HadCM‐driven simulation, however, extreme responses are shown in some regions: Deciduous forest is replaced by grasses in large areas in the middle latitudes, and substantial areas in northern South America and southern Africa predominantly covered by evergreen forest are replaced with grasses while NPP and fire emissions reduce drastically (by more than 100%). A future paper will examine how the biosphere response documented here influences the impact of climate change on surface hydrological conditions.

Water source partitioning among trees growing on shallow karst soils in a seasonally dry tropical climate

The sources of water used by woody vegetation growing on karst soils in seasonally dry tropical regions are little known. In northern Yucatan (Mexico), trees withstand 4-6 months of annual drought in spite of the small water storage capacity of the shallow karst soil. We hypothesized that adult evergreen trees in Yucatan tap the aquifer for a reliable supply of water during the prolonged dry season. The naturally occurring concentration gradients in oxygen and hydrogen stable isotopes in soil, bedrock, groundwater and plant stem water were used to determine the sources of water used by native evergreen and drought-deciduous tree species. While the trees studied grew over a permanent water table (9-20 m depth), pit excavation showed that roots were largely restricted to the upper 2 m of the soil/bedrock profile. At the peak of the dry season, the delta(18)O signatures of potential water sources for the vegetation ranged from 4.1 +/- 1.1 per thousand in topsoil to -4.3 +/- 0.1 per thousand in groundwater. The delta(18)O values of tree stem water ranged from -2.8 +/- 0.3 per thousand in Talisia olivaeformis to 0.8 +/- 1 per thousand in Ficus cotinifolia, demonstrating vertical partitioning of soil/bedrock water among tree species. Stem water delta(18)O values were significantly different from that of groundwater for all the tree species investigated. Stem water samples plotted to the right of the meteoric water line, indicating utilization of water sources subject to evaporative isotopic enrichment. Foliar delta(13)C in adult trees varied widely among species, ranging from -25.3 +/- 0.3 per thousand in Enterolobium cyclocarpum to -28.7 +/- 0.4 per thousand in T. olivaeformis. Contrary to initial expectations, data indicate that native trees growing on shallow karst soils in northern Yucatan use little or no groundwater and depend mostly on water stored within the upper 2-3 m of the soil/bedrock profile. Water storage in subsurface soil-filled cavities and in the porous limestone bedrock is apparently sufficient to sustain adult evergreen trees throughout the pronounced dry season.

Dieback and Episodic Mortality of Cercidium microphyllum (Foothill Paloverde), a Dominant Sonoran Desert Tree

BOWERS, J. E. AND R. M. TURNER. (U. S. Geological Survey, Tucson, AZ 85745). Dieback and episodic mortality of Cercidium microphyllum (foothill paloverde), a dominant Sonoran Desert tree. J. Torrey Bot. Soc. 128:000-000. 2001.-Past and current dieback of Cercidium microphyllum, a dominant, drought-deciduous tree in the Sonoran Desert, was investigated at Tumamoc Hill, Tucson, Arizona, USA. Logistic regression predicted that the odds of a Cercidium plant being alive should decrease with increasing circumference, association with the columnar cactus Carnegiea gigantea, and occurrence on steep slopes. Slope azimuth, parasitization by Phoradendron californicum, and distance to nearest Cercidium within 5 m did not significantly affect the odds of survival. Carnegiea was a source of background mortality rather than a primary cause of dieback. Of the >1,000 living and dead plants sampled, 7.7% had died within the past 5 to 7 years. An additional 12.8% died in the more distant past. Diebacks tended to occur during severe deficits in annual, especially summer, rain. More than half of the dead plants in the sample were 250 cm in girth. In current and past diebacks on Tumamoc Hill, it seems likely that severe drought interacted with natural senescence of an aging population, weakening large, old trees and hastening their deaths.

Ecophysiological studies of Sonoran Desert plants : IV. Seasonal photosynthetic capacities of Acacia greggii and Cercidium microphyllum.

The gas exchange and water relations of two Sonoran Desert plants were measured throughout a 12 month period. Seasonal photosynthesis patterns of both species followed the seasonal variation in soil and plant water potential. Acacia greggii, a winter deciduous shrub, appears to be fall active since the day-long mean photosynthesis rate was maximum, i.e., 16 mg CO2 dm-2 h-1, at this time. Cercidium microphyllum, a drought deciduous and chlorophyllous-stemmed tree, also appears to be mainly fall active. For this species the day-long mean photosynthesis rate was not in excess of 14 mg CO2 dm-2 h-1. Both species initiate leaf production in the spring and neither experiences severe plant water stress. Seasonal minima of dawn plant water potential were-44 and-32 bars for the shrub and tree species, respectively. The two species differ slightly in their tolerance of heat and water stress, since foliated plants of Acacia greggii maintain summer gross photosynthesis.All of the aboveground organs for plants of C. microphyllum are capable of exogenous 14CO2 assimilation. This species appears to be unique in the magnitude of the photosynthetic production contributed by stems. Seasonal production by stems, leaves and flowers/fruits averaged 72, 24 and 4% of the total carbon gain per tree, respectively. Aboveground gross primary production was over 4.5-fold greater than aboveground net primary production. This difference between these two production estimates is likely due to the very small foliar biomass maintained throughout the year and the energy expenditure required to maintain the metabolically active cells of the chlorophyllous stems.

Carbohydrate Storage Cycles in Two Californian Mediterranean-Climate Trees

Summary The annual carbohydrate economy of two trees in the mediterranean climate of California was studied. One, a drought-deciduous tree, Aesculus californica, stores large quantities of carbohydrate during the spring and early summer photosynthetic period which is subsequently used for massive fruit production in fall and for at least a portion of regrowth in February. In contrast, the evergreen oak, Quercus agrifolia, has little annual change in the carbohydrate status of its tissues and presumably channels photosynthate directly into fruits during this long development period. However, some reserves may be involved.